The ecology of fear and inverted biomass pyramids

In inverted biomass pyramids (IBPs) prey are outnumbered by their predators when measured by biomass. We investigate how prey should behave in the face of danger from higher predator biomass, and how anti-predator behavior (in the form of vigilance) can, in turn, affect the predator–prey system. In this study, we incorporate anti-predator behaviors into a Lotka–Volterra predator–prey model in the form of fixed and facultative vigilance. Facultative vigilance models behavior as a dynamic foraging game, allowing us to assess optimal behavioral responses in the context of IBPs using a dynamical fitness optimization approach. We model vigilance as a tradeoff between safety and either the prey's maximum growth rate or its carrying capacity. We assess the population dynamics of predators and prey with fear responses, and investigate the role fear plays on trophic structure. We found that the ecology of fear plays an important role in predator–prey systems, impacting trophic structure and the occurrence of IBPs. Fixed vigilance works against IBP structure by always reducing the predator–prey biomass ratio at equilibrium with increasing levels of vigilance. Facultative vigilance can actually promote IBPs, as prey can now adjust their vigilance levels to cope with increased predation and the costs associated with vigilance. This is especially true when the effectiveness of vigilance is low and predators are very lethal. In general, these trends are true whether the costs of vigilance are felt on the prey's maximum growth rate or its carrying capacity. Just as the ecology of fear, when first introduced, was used to explain why top carnivores are rare in terrestrial systems, it can also be used to understand how big fierce predators can be common in IBPs.     

Controlling chaos in ecology: From deterministic to individual-based models

The possibility of chaos control in biological systems has been stimulated by recent advances in the study of heart and brain tissue dynamics. More recently, some authors have conjectured that such a method might be applied to population dynamics and even play a nontrivial evolutionary role in ecology. In this paper we explore this idea by means of both mathematical and individual-based simulation models. Because of the intrinsic noise linked to individual behavior, controlling a noisy system becomes more difficult but, as shown here, it is a feasible task allowed to be experimentally tested.     

Ecological rigidity and the hardness of selection in the wild

Hutchinson's ecological theater and evolutionary play is a classical view of evolutionary ecology—ecology provides a template in which evolution occurs. An opposing view is that ecological and evolutionary changes are like two actors on a stage, intertwined by density and frequency dependence. These opposing views correspond to hard and soft selection, respectively. Although often presented as diametrically opposed, both types of selection can occur simultaneously, yet we largely lack knowledge of the relative importance of hard versus soft selection in the wild. I use a dataset of 3000 individual gall makers from 15 wild local populations over 5 years to investigate the hardness of selection. I show that enemy attack consistently favors some gall sizes over others (hard selection) but that these biases can be fine-tuned by density and frequency dependence (soft selection). As a result, selection is hard and soft in roughly equal measures, but the importance of each type varies as species interactions shift. I conclude that eco-evolutionary dynamics should occur when a mix of hard and soft selection acts on a population. This work contributes to the rapprochement of disparate views of evolutionary ecology—ecology is neither a rigid theater nor a flexible actor, but instead embodies components of both.     

An acarine predator-prey metapopulation system inhabiting greenhouse cucumbers

The two-spotted spider mite ( Tetranychus urticae ) is a serious pest on greenhouse cucumbers, but can be controlled by the phytoseiid predator Phytoseiulus persimilis. The two mite species exhibit considerable fluctuations in overall population densities but within acceptable limits. The system appears to be persistent at a regional (greenhouse) scale in spite of frequent local extinctions (e.g. at individual plants). Experimental evidence indicates that the mites form a metapopulation system characterized by 'shifting mosaic' dynamics. A stochastic simulation model is used to analyse the role played by dispersal in the dynamics and persistence of the system. It shows that demographic stochasticity generates sufficient endogenous 'noise' to counteract the synchronizing effect of density-dependent dispersal, provided dispersal rates are not too high and the system is not too small. Low dispersal rates, on the other hand, increase the risk of local outbreaks of spider mites that may cause destruction of plants.     

An ecological perspective on marine reserves in prey–predator dynamics

This paper describes a prey–predator type fishery model with prey dispersal in a two-patch environment, one of which is a free fishing zone and other is a protected zone. The existence of possible steady states, along with their local stability, is discussed. A geometric approach is used to derive the sufficient conditions for global stability of the system at the positive equilibrium. Relative size of the reserve is considered as control in order to study optimal sustainable yield policy. Subsequently, the optimal system is derived and then solved numerically using an iterative method with Runge–Kutta fourth-order scheme. Numerical simulations are carried out to illustrate the importance of marine reserve in fisheries management. It is noted that the marine protected area enables us to protect and restore multi-species ecosystem. The results illustrate that dynamics of the system is extremely interesting if simultaneous effects of a regulatory mechanism like marine reserve is coupled with harvesting effort. It is observed that the migration of the resource, from protected area to unprotected area and vice versa, is playing an important role towards the standing stock assessment in both the areas which ultimately control the harvesting efficiency and enhance the fishing stock up to some extent.     

Impacts of predator-mediated interactions along a climatic gradient on the population dynamics of an alpine bird

According to classic theory, species'' population dynamics and distributions are less influenced by species interactions under harsh climatic conditions compared to under more benign climatic conditions. In alpine and boreal ecosystems in Fennoscandia, the cyclic dynamics of rodents strongly affect many other species, including ground-nesting birds such as ptarmigan. According to the ‘alternative prey hypothesis’ (APH), the densities of ground-nesting birds and rodents are positively associated due to predator–prey dynamics and prey-switching. However, it remains unclear how the strength of these predator-mediated interactions change along a climatic harshness gradient in comparison with the effects of climatic variation. We built a hierarchical Bayesian model to estimate the sensitivity of ptarmigan populations to interannual variation in climate and rodent occurrence across Norway during 2007–2017. Ptarmigan abundance was positively linked with rodent occurrence, consistent with the APH. Moreover, we found that the link between ptarmigan abundance and rodent dynamics was strongest in colder regions. Our study highlights how species interactions play an important role in population dynamics of species at high latitudes and suggests that they can become even more important in the most climatically harsh regions.     

Biological control in a disturbed environment

Most ecological and epidemiological models describe systems with continuous uninterrupted interactions between populations. Many systems, though, have ecological disturbances, such as those associated with planting and harvesting of a seasonal crop. In this paper, we introduce host–parasite–hyperparasite systems as models of biological control in a disturbed environment, where the host–parasite interactions are discontinuous. One model is a parasite–hyperparasite system designed to capture the essence of biological control and the other is a host–parasite–hyperparasite system that incorporates many more features of the population dynamics. Two types of discontinuity are included in the models. One corresponds to a pulse of new parasites at harvest and the other reflects the discontinuous presence of the host due to planting and harvesting. Such discontinuities are characteristic of many ecosystems involving parasitism or other interactions with an annual host. The models are tested against data from an experiment investigating the persistent biological control of the fungal plant parasite of lettuce Sclerotinia minor by the fungal hyperparasite Sporidesmium sclerotivorum, over successive crops. Using a combination of mathematical analysis, model fitting and parameter estimation, the factors that contribute the observed persistence of the parasite are examined. Analytical results show that repeated planting and harvesting of the host allows the parasite to persist by maintaining a quantity of host tissue in the system on which the parasite can reproduce. When the host dynamics are not included explicitly in the model, we demonstrate that homogeneous mixing fails to predict the persistence of the parasite population, while incorporating spatial heterogeneity by allowing for heterogeneous mixing prevents fade-out. Including the host''s dynamics lessens the effect of heterogeneous mixing on persistence, though the predicted values for the parasite population are closer to the observed values. An alternative hypothesis for persistence involving a stepped change in rates of infection is also tested and model fitting is used to show that changes in some environmental conditions may contribute to parasite persistence. The importance of disturbances and periodic forcing in models for interacting populations is discussed.     

中国兽类种群生态学研究进展与展望

兽类种群生态学是现代生态学的核心研究内容。Charles Sutherland Elton在20世纪20年代发现小哺乳动物种群波动现象,标志着现代种群生态学研究的开始。什么因素调节种群波动的问题一直是现代种群生态学领域的研究热点。我国兽类种群生态学研究始于20世纪50年代,迄今,已走过了70年的发展历程,并取得了重要成果。本综述基于20世纪50年代以来我国学者在主流中文期刊及科学引文索引（Science Citation Index,简称SCI）刊物发表的历史文献,分别从种群波动格局、种群统计参数变化、种群内部和外部调节等不同层面评述了我国在鼠类和大型兽类种群生态学的研究历程及现状,同时探讨了未来的研究方向。     

On harvesting a structured ungulate population

Variation in demographic rates within a spatially structured population could have important consequences for management decisions, harvesting strategies and offtake rates. Although there is a growing body of evidence suggesting that demographic rates vary within populations over a range of spatial scales, there has been little research investigating the consequences of this variation for population management. In this paper, data on the dynamics of two female red deer sub-populations on Rum are analysed, and evidence is presented for differences between the fecundity and mortality rates of the two sub-populations. A simple harvesting model is developed to represent the dynamics of the two sub-populations, including density-independent migration between sub-populations and spatially correlated environmental variability. The highest monetary yield in the model is obtained by harvesting the more resilient sub-population at a higher rate. Surprisingly the losses involved in harvesting both sub-populations at the same rate are insignificant. However, if migration were density-dependent, the size of one sub-population would be more relevant to harvesting policy for the other sub-population. The results of this empirical study are compared to theoretical work on spatially structured populations; it is shown that when a species has complex age- and sex-structured population dynamics, previous theoretical results may not hold.     

Dynamics of additional food provided predator–prey system with mutually interfering predators

Use of additional/alternative food source to predators is one of the widely recognised practices in the field of biological control. Both theoretical and experimental works point out that quality and quantity of additional food play a vital role in the controllability of the pest. Theoretical studies carried out previously in this direction indicate that incorporating mutual interference between predators can stabilise the system. Experimental evidence also point out that mutual interference between predators can affect the outcome of the biological control programs. In this article dynamics of additional food provided predator–prey system in the presence of mutual interference between predators has been studied. The mutual interference between predators is modelled using Beddington–DeAngelis type functional response. The system analysis highlights the role of mutual interference on the success of biological control programs when predators are provided with additional food. The model results indicate the possibility of stable coexistence of predators with low prey population levels. This is in contrast to classical predator–prey models wherein this stable co-existence at low prey population levels is not possible. This study classifies the characteristics of biological control agents and additional food (of suitable quality and quantity), permitting the eco-managers to enhance the success rate of biological control programs.     

黄土高原集水农业研究进展

回顾了黄土高原集水农业理论与技术体系的研究成果．分析评价了集水农业的研究进展。随着黄土高原集水农业研究方法的改进、研究内容的深入、研究领域的扩充．提出了广义性集水农业研究范畴。在黄土高原集水农业理论研究的基础上．应加强微集雨微灌溉应用技术、现代集雨技术、计算机控制技术与集雨网络等高新技术手段的技术集成．以提高雨水汇集与利用效率。同时．黄土高原集水农业的研究已经从微生境条件下的农业生态系统延伸至区域生态环境保育。利用汇集雨水合理调配生态用水．进行小流域综合治理。农林牧综合发展。生态环境重建的集水型生态农业是黄土高原集水农业的发展趋势。     

The effect of seasonal harvesting on stage-structured population models

In most models of population dynamics, increases in population due to birth are assumed to be time-independent, but many species reproduce only during a single period of the year. We propose an exploited single-species model with stage structure for the dynamics in a fish population for which births occur in a single pulse once per time period. Since birth pulse populations are often characterized with a discrete time dynamical system determined by its Poincaré map, we explore the consequences of harvest timing to equilibrium population sizes under seasonal dependence and obtain threshold conditions for their stability, and show that the timing of harvesting has a strong impact on the persistence of the fish population, on the volume of mature fish stock and on the maximum annual-sustainable yield. Moreover, our results imply that the population can sustain much higher harvest rates if the mature fish is removed as early in the season (after the birth pulse) as possible. Further, the effects of harvesting effort and harvest timing on the dynamical complexity are also investigated. Bifurcation diagrams are constructed with the birth rate (or harvesting effort or harvest timing) as the bifurcation parameter, and these are observed to display rich structure, including chaotic bands with periodic windows, pitch-fork and tangent bifurcations, non-unique dynamics (meaning that several attractors coexist) and attractor crisis. This suggests that birth pulse, in effect, provides a natural period or cyclicity that makes the dynamical behavior more complex.This work is supported by National Natural Science Foundation of China (10171106)     

Adaptive Control Based Harvesting Strategy for a Predator–Prey Dynamical System

This paper deals with designing a harvesting control strategy for a predator–prey dynamical system, with parametric uncertainties and exogenous disturbances. A feedback control law for the harvesting rate of the predator is formulated such that the population dynamics is asymptotically stabilized at a positive operating point, while maintaining a positive, steady state harvesting rate. The hierarchical block strict feedback structure of the dynamics is exploited in designing a backstepping control law, based on Lyapunov theory. In order to account for unknown parameters, an adaptive control strategy has been proposed in which the control law depends on an adaptive variable which tracks the unknown parameter. Further, a switching component has been incorporated to robustify the control performance against bounded disturbances. Proofs have been provided to show that the proposed adaptive control strategy ensures asymptotic stability of the dynamics at a desired operating point, as well as exact parameter learning in the disturbance-free case and learning with bounded error in the disturbance prone case. The dynamics, with uncertainty in the death rate of the predator, subjected to a bounded disturbance has been simulated with the proposed control strategy.     

Differences in foraging ecology determine variation in visual fields in ibises and spoonbills (Threskiornithidae)

Variations in visual field topography among birds have been interpreted as adaptations to the specific perceptual challenges posed by the species’ foraging ecology. To test this hypothesis we determined visual field topography in four bird species which have different foraging ecologies but are from the same family: Puna Ibis Plegadis ridgwayi (probes for prey in the soft substrates of marsh habitats), Northern Bald Ibis Geronticus eremita (surface pecks for prey in dry terrestrial habitats), African Spoonbill Platalea alba and Eurasian Spoonbill Platalea leucorodia (bill‐sweeps for prey in shallow turbid waters). All four species employ tactile cues provided by bill‐tip organs for prey detection. We predicted that the visual fields of these species would show general features similar to those found in other birds whose foraging is guided by tactile cues from the bill (i.e. bill falling outside the frontal binocular field and comprehensive visual coverage of the celestial hemisphere). However, the visual fields of all four species showed general features characteristic of birds that take food directly in the bill under visual guidance (i.e. a narrow and vertically long binocular field in which the projection of the bill tip is approximately central and with a blind area above and behind the head). Visual fields of the two spoonbills were very similar but differed from those of the ibises, which also differed between themselves. In the spoonbills, there was a blind area below the bill produced by the enlarged spatulate bill tip. We discuss how these differences in visual fields are related to the perceptual challenges of these birds’ different foraging ecologies, including the detection, identification and ingestion of prey. In particular we suggest that all species need to see binocularly around the bill and between the opened mandibles for the identification of caught prey items and its transport to the back of the mouth. Our findings support the hypothesis that sensory challenges associated with differences in foraging ecology, rather than shared ancestry or the control of locomotion, are the main determinants of variation in visual field topography in birds.     

基于管理目标的湖泊生态系统动力学

基于生态系统管理的目标,在对相关研究分析的基础上,依据生态系统生态学、淡水生态学的理论,提出了湖泊生态系统动力学研究的2个理论基础：生态系统管理和生态系统特征.在此基础上,分析得到湖泊生态系统动力学的研究方法体系,主要包括研究内容与技术路线、关键问题识别和动力学模拟、湖泊生态系统的适应性管理决策等部分.其中,湖泊生态系统结构和过程、湖泊中食物网营养动力学研究、生源要素循环、湖泊中关键过程的生态作用以及湖泊生态系统动力学模拟是研究的核心问题.此后,以P为主要的生源要素,将生态系统分为3个子过程：入流、出流和内部反馈,并以此建立了湖泊生态系统动力学的模型框架,以辅助于湖泊的生态系统管理.     

Bottom‐up processes drive reproductive success in an apex predator

One of the central goals of the field of population ecology is to identify the drivers of population dynamics, particularly in the context of predator–prey relationships. Understanding the relative role of top‐down versus bottom‐up drivers is of particular interest in understanding ecosystem dynamics. Our goal was to explore predator–prey relationships in a boreal ecosystem in interior Alaska through the use of multispecies long‐term monitoring data. We used 29 years of field data and a dynamic multistate site occupancy modeling approach to explore the trophic relationships between an apex predator, the golden eagle, and cyclic populations of the two primary prey species available to eagles early in the breeding season, snowshoe hare and willow ptarmigan. We found that golden eagle reproductive success was reliant on prey numbers, but also responded prior to changes in the phase of the snowshoe hare population cycle and failed to respond to variation in hare cycle amplitude. There was no lagged response to ptarmigan populations, and ptarmigan populations recovered quickly from the low phase. Together, these results suggested that eagle reproduction is largely driven by bottom‐up processes, with little evidence of top‐down control of either ptarmigan or hare populations. Although the relationship between golden eagle reproductive success and prey abundance had been previously established, here we established prey populations are likely driving eagle dynamics through bottom‐up processes. The key to this insight was our focus on golden eagle reproductive parameters rather than overall abundance. Although our inference is limited to the golden eagle–hare–ptarmigan relationships we studied, our results suggest caution in interpreting predator–prey abundance patterns among other species as strong evidence for top‐down control.     

Testing the validity of functional response models using molecular gut content analysis for prey choice in soil predators

Analysis of predator–prey interactions is a core concept of animal ecology, explaining structure and dynamics of animal food webs. Measuring the functional response, i.e. the intake rate of a consumer as a function of prey density, is a powerful method to predict the strength of trophic links and assess motives of prey choice, particularly in arthropod communities. However, due to their reductionist set‐up, functional responses, which are based on laboratory feeding experiments, may not display field conditions, possibly leading to skewed results. Here, we tested the validity of functional responses of centipede predators and their prey by comparing them with empirical gut content data from field‐collected predators. Our predator–prey system included lithobiid and geophilomorph centipedes, abundant and widespread predators of forest soils and their soil‐dwelling prey. First, we calculated the body size‐dependent functional responses of centipedes using a published functional response model in which we included natural prey abundances and animal body masses. This allowed us to calculate relative proportions of specific prey taxa in the centipede diet. In a second step, we screened field‐collected centipedes for DNA of eight abundant soil‐living prey taxa and estimated their body size‐dependent proportion of feeding events. We subsequently compared empirical data for each of the eight prey taxa, on proportional feeding events with functional response‐derived data on prey proportions expected in the gut, showing that both approaches significantly correlate in five out of eight predator–prey links for lithobiid centipedes but only in one case for geophilomorph centipedes. Our findings suggest that purely allometric functional response models, which are based on predator–prey body size ratios are too simple to explain predator–prey interactions in a complex system such as soil. We therefore stress that specific prey traits, such as defence mechanisms, must be considered for accurate predictions.     

From interesting details to dynamical relevance: toward more effective use of empirical insights in theory construction

A perennial challenge in ecology is to develop dynamical systems models that appropriately abstract and characterize the dynamics of natural systems. Deriving an appropriate model of system dynamics can be a long and iterative process whose outcome depends critically on the quality of empirical data describing the long‐term behavior of a natural system. Most ecological time series are insufficient to offer insight into the way organizational hierarchies and spatial scales are causally linked to natural system dynamics. Moreover, the classic tradition of hypothesis testing in ecology is not likely to lead to those key insights. This because empirical research is geared almost exclusively toward testing model predictions based on underlying causal relationships assumed by theorists. So, empirical research relies heavily on theory for guidance on what is or is not dynamically relevant. I argue here that it is entirely possible to reduce much of this guesswork involved with deciding on causal structure by giving empirical research a new role in theory development. In this role, natural history and field observations are used to develop stochastic, individual‐based and spatially explicit computational models or IBMs that can explore the range of contingency and complexity inherent in real‐world systems.
IBMs can be used to run simulations allowing deductions to be made about the causal linkages between organizational hierarchies, spatial scales, and dynamics. These deductions can be tested under field conditions using experiments that manipulate the putative causal structure and evaluate the dynamical consequences. The emerging insights from this stage can then be used to inspire an analytical construct that embodies the dynamically relevant scales and mechanisms. In essence, computational modeling serves as an intermediate step in theory development in that a wide range of possibly important biological details are considered and then reduced to a subset that is dynamically relevant.     

Integrating terrestrial scavenging ecology into contemporary wildlife conservation and management

Scavenging plays a vital role in maintaining ecosystem health and contributing to ecological functions; however, research in this sub‐discipline of ecology is underutilized in developing and implementing wildlife conservation and management strategies. We provide an examination of the literature and recommend priorities for research where improved understanding of scavenging dynamics can facilitate the development and refinement of applied wildlife conservation and management strategies. Due to the application of scavenging research broadly within ecology, scavenging studies should be implemented for informing management decisions. In particular, a more direct link should be established between scavenging dynamics and applied management programs related to informing pharmaceutical delivery and population control through bait uptake for scavenging species, prevention of unintentional poisoning of nontarget scavenging species, the epidemiological role that scavenging species play in disease dynamics, estimating wildlife mortalities, nutrient transfer facilitated by scavenging activity, and conservation of imperiled facultative scavenging species. This commentary is intended to provide information on the paucity of data in scavenging research and present recommendations for further studies that can inform decisions in wildlife conservation and management. Additionally, we provide a framework for decision‐making when determining how to apply scavenging ecology research for management practices and policies. Due to the implications that scavenging species have on ecosystem health, and their overall global decline as a result of anthropic activities, it is imperative to advance studies in the field of scavenging ecology that can inform applied conservation and management programs.